© 2005 Nature Publishing Group
T
he announcement by Kasahara and Kato
of a new redox-cofactor vitamin for
mammals
1
, pyrroloquinoline quinone
(PQQ), was based on their claim that an
enzyme, predicted to be involved in mouse
lysine metabolism, is a PQQ-dependent
dehydrogenase. However, this claim was
dependent on a sequence analysis using data-
bases that inappropriately label ȋ-propeller
sequences as PQQ-binding motifs. What
the evidence actually suggests is that the
enzyme is an interesting novel protein that
has a seven-bladed ȋ-propeller structure,
but there is nothing to indicate that it is a
PQQ-dependent dehydrogenase.
In bacteria, PQQ is an essential cofactor
for various dehydrogenase enzymes known
as quinoproteins
2
. Nutritional experiments
have indicated some (unknown) metabolic
or nutritional role for PQQ in mammals
1,3,4
,
but it cannot be accepted as a vitamin until it
is proved to be required by an enzyme as an
essential cofactor; this is the key evidence that
Kasahara and Kato
1
claim to have provided.
In the course of a study on bipolar disorder
(see www.brain.riken.go.jp/labs/mdmd/pqq/
index-e.html), these authors cloned a mouse
gene encoding a protein (U26) with some
similarity to yeast aminoadipate reductase
(LYS2)
5
; they proposed that mouse U26
could be involved in one of the important
first steps in the degradation of dietary
lysine, acting as a PQQ-dependent adipic 6-
semialdehyde dehydrogenase.
As would be expected from the method
used by Kasahara and Kato in searching for
the LYS2 analogue
1
, the U26 sequence con-
tained no carboxy-terminal NAD(P)-bind-
ing domain. They noted from sequence
analysis that U26 has an alternative carboxy-
terminal domain that contains seven repeats
of the ‘PQQ-binding motif ’ that is conserved
among bacterial PQQ-dependent dehydro-
genase enzymes, leading to the conclusion
that mouse U26 could be a PQQ-dependent
dehydrogenase.
However, this conclusion is based on the
misconception that the Smart and Pfam
brief communications arising
NATURE | VOL 433 | 27 JANUARY 2005 | www.nature.com/nature E10
databases are able to recognize PQQ-bind-
ing sites even when, as in this case, there is
negligible sequence similarity to known
PQQ-dependent enzymes. The ‘sites’ wrongly
identified by the databases do not represent
PQQ-binding sites but represent the ȋ-
sheets that form the ‘blades’ of the ‘propeller
fold’ that happens to be a feature of all PQQ-
dependent dehydrogenases, whose main
structure is a superbarrel made up of either
six or eight ‘propeller blades’ (Fig. 1). The
propeller fold is not related in any direct
way to PQQ binding
2,6
, and these folds are
found in many other types of protein,
which have extreme functional and phylo-
genetic diversity
7
.
We contend that there is still no com-
pelling evidence for a PQQ-dependent
enzyme in the mouse and that the authors’
announcement of a new vitamin was there-
fore premature.
Leigh M. Felton, Chris Anthony
School of Biological Sciences, University of
Southampton, Southampton SO16 7PX, UK
e-mail: c.anthony@soton.ac.uk
doi:10.1038/nature03322
1. Kasahara, T. & Kato, T. Nature 422, 832 (2003).
2. Goodwin, P. M. & Anthony, C. Adv. Microb. Physiol. 40, 1–80
(1998).
3. McIntire, W. S. Annu. Rev. Nutr. 18, 145–177 (1998).
4. Steinberg, F. et al. Exp. Biol. Med. 228, 160–166 (2003).
5. Ehmann, D. E., Gehring, A. M. & Walsh, C. T. Biochemistry 38,
6171–6177 (1999).
6. Anthony, C. & Ghosh, M. Progr. Biophys. Mol. Biol. 69, 1–21
(1998).
7. Paoli, M. Progr. Biophys. Mol. Biol. 76, 103–130 (2001).
Reply: T. Kasahara and T. Kato reply to this communication
(doi:10.1038/nature03324).
Biochemistry
Is pyrroloquinoline
quinone a vitamin?
T
he announcement by Kasahara and
Kato
1
of pyrroloquinoline quinone
(PQQ) as a ‘new’ vitamin has received
considerable attention. We have since
attempted to reproduce the findings on
which their conclusion is based, namely
that defects in lysine metabolism occur in
PQQ-deprived rodents. However, we find
that the activity of Ȋ-aminoadipic acid-Ȏ-
semialdehyde (AAS) dehydrogenase in liver
and plasma levels of Ȋ-aminoadipic acid
(AAA), both of which act as indicators of
lysine degradation in mammals, are not
affected by changes in PQQ dietary status.
Our results call into question the identifica-
tion of PQQ as a new vitamin.
The observations of Kasahara and Kato
1
were of particular interest to us because their
experimental model was based on our find-
ings that mice that are fed chemically defined
diets and deprived of PQQ often show signs
of reproductive failure and compromised
neonatal growth
2,3
. Using previously
described assay conditions
4
, we found that
Biochemistry
Role of PQQ as a mammalian enzyme cofactor?
Arising from: T. Kasahara & T. Kato Nature 422, 832 (2003)
Figure 1 The superbarrel structure of the Ȋ-subunit of the PQQ-dependent methanol dehydrogenase
6
. This is simplified to show the ȋ-
sheets (labelled W1–W8) that form the ‘propeller blades’ of the propeller fold; each blade is made up of four antiparallel strands (labelled
A–D). It can be seen that these eight ȋ-sheets have no specific role in binding the PQQ molecule (shown as ball-and-stick model). The
calcium ion is shown as a small sphere.